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Open AcceShort ReportTwo new endemic genera and a new species of toad (Anura: Bufonidae) from the Western Ghats of IndiaSD Biju1, Ines Van Bocxlaer2, Varad B Giri3, Simon P Loader4 and Franky Bossuyt*2

Address: 1Systematics Lab, Centre for Environmental Management of Degraded Ecosystems (CEMDE), School of Environmental Studies, University of Delhi 110 007, India, 2Biology Department, Unit of Ecology & Systematics: Amphibian Evolution Lab, Vrije Universiteit Brussel (VUB), Pleinlaan 2, B-1050 Brussels, Belgium, 3Collections Department, Bombay Natural History Society (BNHS), S.B. Singh Road, Mumbai 400 001, India and 4Institute of Biogeography, Department of Environmental Sciences, University of Basel, Klingelbergstrasse 27, 4056 Basel, Switzerland

Email: SD Biju - sdbiju@cemde.du.ac.in; Ines Van Bocxlaer - ivbocxla@vub.ac.be; Varad B Giri - varadgiri@gmail.com; Simon P Loader - Simon.Loader@unibas.ch; Franky Bossuyt* - fbossuyt@vub.ac.be

* Corresponding author

AbstractBackground: Bufonidae are a large family of toads with a subcosmopolitan distribution. Recentmolecular phylogenetic analyses have revealed a radiation of toads (Adenominae) with distinct adultand larval ecomorphs on the Southern parts of the Indian subcontinent. The Indian torrentialspecies "Ansonia" ornata has a basal position in this clade and does not group with South-East AsianAnsonia. Additionally, the nested position of "Bufo" koynayensis and an undescribed sister species,and their distinct ecologies including a non-typical egg-laying strategy within bufonids, support therecognition of a second distinct genus. In this paper we describe two new genera and one newspecies from the Adenominae clade.

Findings: Ansonia ornata Günther, 1876 "1875" is transferred to Ghatophryne gen. nov., a genusof torrentially adapted toads that are endemic to the Western Ghats of India. On the basis of closemorphological resemblance and distribution, Ansonia rubigina Pillai and Pattabiraman, 1981 isprovisionally transferred to this new genus. The Western Ghats endemic toad Bufo koynayensisSoman, 1963 is transferred to a new genus Xanthophryne gen. nov. Based on molecular andmorphological evidence, we additionally describe a new species, Xanthophryne tigerinus sp. nov.,from Amboli in the Western Ghats.

Conclusion: The descriptions and subsequent taxonomic changes we propose result in threegenera of bufonids recognised as being endemic to the Western Ghats (Ghatophryne gen. nov.,Xanthophryne gen. nov. and Pedostibes), and one to Sri Lanka (Adenomus). The spatialdistribution, and arrangement of these lineages at the base of Adenominae diversification, reflectstheir Early Neogene isolation in the Western Ghats-Sri Lanka hotspot.

Published: 7 December 2009

BMC Research Notes 2009, 2:241 doi:10.1186/1756-0500-2-241

Received: 7 May 2009Accepted: 7 December 2009

This article is available from: http://www.biomedcentral.com/1756-0500/2/241

© 2009 Bossuyt et al; licensee BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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IntroductionBufonidae are a family of toads with over 500 extant spe-cies distributed on most continents. Current opinions onbufonid taxonomy are very divergent, and range from therecognition of multiple genera [1], to favouring a subcos-mopolitan genus Bufo with plenty of subgenera [2].Although the phylogenetic relationships of bufonids havebeen studied intensively [1,3,4], the evolutionary positionof species on the Indian subcontinent had remainedunclear. However, recent molecular phylogenetic analysesrevealed that these toads belong to a radiation containingdistinct ecomorphs in adult and/or larval forms [5]. Theearly diversification of this clade, for which the name Ade-nominae is available [6], has been reconstructed as occur-ring on the Southern parts of the Indian subcontinent.Early diversification of Adenominae, timed at the earlyMiocene, led to several endemic lineages ("Ansonia"ornata, Pedostibes tuberculosus, the "Bufo" koynayensis groupand Adenomus) [5] and to the origin of the more wide-spread Duttaphrynus clade. The unexpected evolutionaryposition of "Ansonia" ornata and "Bufo" koynayensis (seeabstract) requires recognition of two new genera toaccommodate these Western Ghats endemic species andtheir relatives.

MethodsCollection of amphibian specimens was made duringfieldtrips in the Western Ghats between 1997 and 2002.Specimens were preserved in 5% formaldehyde for 2 days,and subsequently transferred to 70% ethanol. Measure-ments were taken to the nearest 0.1 mm, using a digitalslide-caliper or a binocular microscope with a micrometerocular. The description (all measurements in mm) of thetypes follows terminology used elsewhere [7]. To complywith regulations of the International Code of ZoologicalNomenclature we have deposited copies of this article atthe following publicly accessible libraries: Bombay Natu-ral History Society, Mumbai, India (BNHS); KoninklijkBelgisch Instituut voor Natuurwetenschappen, Brussels,Belgium (KBIN); Natural History Museum, London, UK(BMNH); American Museum of Natural History, NewYork, USA (AMNH); Museum National d'HistoireNaturelle, Paris, France (MNHN); Russian Academy ofSciences, Moscow, Russia (RAS).

Descriptions1. Ghatophryne gen. nov. (Figure 1)EtymologyThe generic epithet is derived from the words 'Ghats' and'phryne'. The former is the Sanskrit word for 'steps' andrefers to the Western Ghats mountain range, where thisgenus is endemic, while 'phryne' means toad in Greek.

Type speciesAnsonia ornata Günther, "1876" 1875, by original designa-tion.

The original description of this species was based on ninesyntypes (BMNH 1947.2.20.65-1947.2.20.73) collectedin "Bramagherries" (= Brahmagiri Hills, Coorg) by R. H.Beddome. We hereby designate one of them, BMNH1947.2.20.66 (ex BMNH 74.4.29.945), an adult male,SVL 29.0, as lectotype of this nominal taxon.

Diagnosis and comparisonGhatophryne can be distinguished from other bufonidgenera by the combination of the following characters:small-sized adults (male SVL 26.9-30.1, N = 5; female35.0, N = 1); dorsum reddish brown, ventrum darkbrownish black with prominent yellowish-orange spots;head without cranial ridge, lack of parotoid glands; fingersfree, toes medium webbed; skin on dorsum with sparselygranular projections especially on anterior half of thebody; eggs non-pigmented, tadpoles with a suctorial disk,adapted to mountain streams throughout the life cycle(SD Biju, personal observations). Ghatophryne can becharacterized in a phylogenetic framework as the mostinclusive clade that contains Ansonia ornata Günther,"1876" 1875, but not Bufo melanostictus Schneider, 1799and Ansonia hanitschi Inger, 1960 [5].

Ghatophryne is morphologically very similar to species ofAnsonia, but is endemic to the southern part of the West-ern Ghats, while Ansonia is known only from South-EastAsia (Sumatra, Borneo and Philippines).

ContentsGhatophryne ornata (Günther, "1876" 1875) comb.nov., Ghatophryne rubigina (Pillai and Pattabiraman,1981) comb. nov. The latter is provisionally includedbased on distributional data and morphological resem-blances [8].

Ghatophryne ornata specimen SDB 6361, adult male, SVL 29.2 mm, collected from Kurichiyarmala, 11°35'N, 75°58'E, Waya-nad, KeralaFigure 1Ghatophryne ornata specimen SDB 6361, adult male, SVL 29.2 mm, collected from Kurichiyarmala, 11°35'N, 75°58'E, Wayanad, Kerala.

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DistributionGhatophryne has only been reported from Kerala and Kar-nataka in the Western Ghats of India.

Description of lectotype of Ansonia ornata Günther, "1876" 1875Medium sized frog (SVL 29.0), body rather elongate.Head slightly longer than wide (HL 8.9; HW 8.5; MN 7.2;MFE 5.4; MBE 1.3); outline of snout in dorsal view nearlyoval, its length (SL 4.5) larger than the horizontal diame-ter of the eye (EL 3.7); loreal region vertically acute; dis-tance between anterior margins of eyes (IFE 4.6) 1.8 timesin distance between posterior margins of eyes (IBE 8.4);interorbital area (IUE 2.8) broader than upper eyelidwidth (UEW 2.3); tympanum (TYD 2.7) rather indistinct;supratympanic fold rather indistinct; parotoid glandsabsent; parietal ridges absent; tongue elongate. Forearm(FLL 7.5) shorter than hand (HAL 8.8; TFL 4.0), strong;relative length of fingers, shortest to longest: I < II < IV <III, tip of fingers enlarged, rounded, without distinctgrooves, without lateral dermal fringe, webbing absent;subarticular tubercles weakly developed; supernumerarytubercles weakly developed; palmar tubercle absent, nup-tial pads present. Hind limbs moderately long, shank(ShL 14.1) longer than thigh (TL 13.3), longer than dis-tance from base of inner metatarsal tubercle to tip of toeIV (FOL 12.6), shorter than distance from heel to tip of toeIV (TFOL 19.2); relative length of toes, shortest to longest:I < II < V < III < IV, tips of toes enlarged, rounded, webbingreduced, reaching up to second subarticular tubercle onboth sides of toe IV; dermal fringe along toe V absent; sub-articular tubercles distinct; supernumerary tuberclesweakly developed; inner metatarsal tubercle elongate(IMT 1.5), outer metatarsal tubercle smaller but distinctlyraised. Skin of snout, between eyes, and upper eyelids sha-greened to sparsely granular, side of head, back, flanksand dorsal part of limbs sparsely granular.

Colour of lectotype: in preservation dorsally uniform lightgreyish brown with a dorsal light yellowish brown patch;ventral side light brown with white irregular spots.

Other specimens studiedOvary of single female specimen (SDB 022) had 35 non-pigmented, creamy white eggs, 1.3 ± 0.4 mm in diameter.Dorsum can have continuous (SDB 6362) or discontinu-ous (SDB 6361) yellowish bands.

2. Xanthophryne gen. novEtymologyderived from two Greek words, 'xanthos' meaning yellow,and 'phryne' meaning toad.

Type speciesBufo koynayensis Soman, 1963, by original designation,ZSIC A1784, Shivaji Sagar lake at Koyna (= Humbarli Vil-lage), Maharashtra, collected by P. W. Soman.

Diagnosis and comparisonXanthophryne can be distinguished from other bufonidgenera by the combination of the following characters:small-sized adults (male SVL 26.5-32.9, N = 12; femaleSVL 33.3-35.3, N = 3) having light brown dorsum with asuffusion of dull chrome-yellow; head with discontinu-ous and weak canthal and preorbital ridges on the ante-rior part, flanks and sides of the abdomen have chrome-yellow patches, or sometimes 2-4 continuous bands; tym-panum indistinct, rather weak parotoid glands; toes andfingers without webbing, tips rounded; eggs in clutches.Xanthophryne can be characterized in a phylogeneticframework as the most inclusive clade that contains Bufokoynayensis Soman, 1963 but not Bufo melanostictusSchneider, 1799 and Bufo kelaartii Günther, 1858.

ContentsXanthophryne koynayensis (Soman, 1963) comb. nov. (Fig-ure 2) and Xanthophryne tigerinus sp. nov. (Figure 3A)

DistributionXanthophryne has only been reported from the northernpart of the Western Ghats of India.

3. Xanthophryne tigerinus sp. nov.(Figure 3A-E)EtymologyThe species epithet tigerinus is attributed to this speciesdue to similarity in the species colour pattern to the prom-inent lateral stripes in Tigers (Panthera tigris).

Type materialHolotype, BNHS 5175, an adult male, SVL 30.5, collectedby SDB and VG on 23 August 2002 from Amboli,15°56'N, 74°00'E, about 720 m asl, Sindhudurg, Mahar-ashtra, India; paratypes, BNHS 5177, an adult male,BNHS 5176, an adult female, collected along with the hol-

Xanthophryne koynayensis, a male adult (SDB 6040) from the type localityFigure 2Xanthophryne koynayensis, a male adult (SDB 6040) from the type locality.

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otype; BNHS 4064, an adult male, BNHS 4063, an adultfemale, collected by VG & Hegde on 9 September 2000,BNHS 4208-4209, two males, BNHS 4207, an adultfemale, collected by Kehimkar and Agarwal on 16 August2003.

Diagnosis and comparisonXanthophryne tigerinus can be distinguished by the follow-ing combination of characters: (1) medium size, maleadult SVL 27.8-32.9, female adult SVL 33.3-35.3; (2) bodyrather elongate; (3) presence of discontinuous canthaland preorbital ridges; (4) stripes on lateral and dorsalside; (5) absence of webbing between fingers and toes.

Xanthophryne tigerinus sp. nov. (Figure 3A-E) differs fromX. koynayensis (Figure 2, 3F) by the presence of a denserarrangement of granular projections with horny spinuleson dorsal and lateral parts of head, back and flank; more

prominent canthal and preorbital ridges; snout longerthan eye length (SL 4.0 ± 0.3 mm, EL 3.4 ± 0.5 mm, N =5, male) vs. snout shorter than eye length (SL 3.3 ± 0.3mm, EL 4.3 ± 0.2 mm, N = 7, male); shank longer thanthigh (ShL 11.5 ± 0.6 mm, TL 9.9 ± 0.4 mm, N = 5, males)vs. shank about equal to thigh (ShL 11.1 ± 0.7 mm, TL11.1 ± 0.8 mm, N = 7, male); foot length longer thanshank and thigh (FOL 12.8 ± 0.6, N = 5, male) vs. equalto shank and thigh (FOL 11.2 ± 0.7, N = 7, male).

Description of the holotypeMedium sized frog (SVL 30.5), body rather elongate (Fig-ure 3A). Head length subequal (HL 10.1) to head width(HW 10.3; MN 8.8; MFE 7.2; MBE 4.2) (Figure 3B, C); out-line of snout in dorsal view nearly oval, in ventral viewoval to pointed, its length (SL 4.1) longer than the hori-zontal diameter of the eye (EL 3.1); loreal region verticallyacute; distance between anterior margins of eyes (IFE 4.6)1.9 times in distance between posterior margins of eyes(IBE 8.7); interorbital area (IUE 2.3) less than upper eye-lid (UEW 3.1); tympanum rather indistinct; supratym-panic fold absent; parotoid glands present, rounded;canthal and preorbital ridges well developed, continuous;tongue entire, oval. Forearm (FLL 6.1) shorter than hand(HAL 7.0; TFL 3.5); relative length of fingers, shortest tolongest: I < II < IV < III (Figure 3D), tip of fingers enlarged,rounded, without distinct grooves, without lateral dermalfringe, webbing absent; subarticular tubercles weaklydeveloped; prepollex rounded, prominent, palmar tuber-cle one, rounded, prominent; supernumerary tuberclesweakly developed; nuptial pads present. Hindlimbs mod-erately long, shank (ShL 11.8) longer than thigh (TL10.1), shorter than distance from base of inner metatarsaltubercle to tip of toe IV (FOL 13.1); distance from heel totip of toe IV (TFOL18.1); relative length of toes, shorter tolongest: I < II < III < V < IV (Figure 3E), tips of toesenlarged, rounded, webbing absent; subarticular tuberclesrather indistinct; supernumerary tubercles weakly devel-oped; inner metatarsal tubercle elongated (IMT 1.2). Skinof snout, between eyes, upper eyelids, side of head, andback have granular projections with horny spinules; dor-sal part of limbs granular with horny spinules; throat sha-greened to granular; chest, belly, and posterior surface ofthighs granular.

Color: in preservation: dorsum light-brown, loreal and tym-panic region light brown, lateral region dark grey withlight colour stripes; fingers I and II, and toes I and IIcreamy white; ventral side light brownish grey with darkgrey irregular spots; in life: dorsum golden yellow, lateralregion brown with light yellow stripes extending fromdorsum; fingers I and II, and toes I and II whitish; ventralside light grey with dark greyish brown irregular spots,lower jaw margins white.

Xanthophryne tigerinus sp. nov (holotype, BNHS 5175)Figure 3Xanthophryne tigerinus sp. nov (holotype, BNHS 5175). A, holotype in life; B, dorsal view of head showing prominent granular projections with horny spinules; C, lat-eral view of head; D, ventral view of hand; E, ventral view of foot; F. Xanthophryne koynayensis, dorsal view of the head, showing scattered granular projections with horny spinules.

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Secondary sexual characteristicsMale (BNHS 5175) has nuptial spines on the prepollex;Ovary of single female specimen ((BNHS 4207) had 59pigmented eggs, 1.2 ± 0.3 mm in diameter.

VariationMeasurements of eight type specimens are given in Table1. During the breeding season, both male and femaletoads have a bright yellow dorsum with light yellow andlight blue lateral stripes. Outside the breeding season,they have a light brownish grey dorsum with light yellowlateral stripes.

Distribution and natural historyXanthophryne tigerinus sp. nov. is known only from thetype locality Amboli. The type series was collected fromthe ground near disturbed evergreen forest patches duringa rainy evening after 19:00 h. Amplexus is axillary. Eggclutches contain 30-35 eggs per clutch (N = 4; VG, per-

sonal observation). The eggs are laid in temporary pud-dles on laterite rocks (Figure 4).

NotesBufo koynayensis Soman, 1963 was described twice basedon the same type series but bearing different numbers.Soman first described Bufo koynayensis, without designat-ing any type other than mentioning, "was collected ingood numbers" [9]. Almost 40 years later, specimen ZSIA.1784 was recognized as the holotype of Bufo koynayensisSoman, 1963 [10]. This specimen was examined and isbadly damaged (SDB, personal observation). The samplescollected by Soman were utilized to describe Bufo sul-phureus (holotype, BNHS 377 and 26 paratypes including5 males, 11 females and 10 juveniles) [11], which as aconsequence is a junior synonym of Bufo koynayensis. Asdescribed in the original description, the type series ofBufo sulphureus were deposited in the British Museum ofNatural History, the Indian Museum and the Bombay

Table 1: Measurements of the two Xanthophryne gen. nov. species

Xanthophryne tigerinus sp. nov. (type series)

Male Female

BNHS5175

BNHS5177

BNHS4064

BNHS 4208

BNHS4209

Mean STDV BNHS5176

BNHS 4063

BNHS 4207

Mean STDV

SVL 30.5 31.0 32.9 32.0 27.8 30.8 1.9 33.9 33.3 35.3 34.2 1.0HW 10.3 10.1 11.0 10.2 10.1 10.3 0.4 12.2 11.5 13.9 12.5 1.2HL 10.1 10.0 10.7 10.1 9.0 10.0 0.6 11.2 10.9 12.2 11.4 0.7IUE 2.3 2.1 2.3 2.1 2.5 2.3 0.2 2.3 2.5 3.0 2.6 0.4

UEW 3.1 2.9 3.6 3.1 2.9 3.1 0.3 3.4 3.4 3.2 3.3 0.1SL 4.1 3.6 4.5 4.1 3.8 4.0 0.3 4.5 4.8 4.6 4.6 0.2EL 3.1 3.0 4.0 4.0 3.0 3.4 0.5 4.4 4.6 4.5 4.5 0.1FLL 6.1 6.8 5.4 6.5 5.8 6.1 0.6 7.4 6.1 6.1 6.5 0.8HAL 7.0 6.9 6.7 6.8 6.7 6.8 0.1 6.7 6.9 7.4 7.0 0.4ShL 11.8 11.9 11.8 11.7 10.4 11.5 0.6 11.6 11.7 12.2 11.8 0.3TL 10.1 10.2 9.3 10.3 9.7 9.9 0.4 10.7 8.8 11.6 10.4 1.4

FOL 13.1 12.8 13.4 12.6 11.9 12.8 0.6 12.6 13.2 13.9 13.2 0.7

Xanthophryne koynayensis

MalesBNHS*

377BNHS**

357BNHS**

358BNHS**

372BNHS**

365BNHS5174

BNHS 5188

Mean STDV

SVL 29.6 29.8 31.8 31.1 31.2 27.1 26.5 29.6 2.1HW 11.1 10.3 11.0 11.0 10.9 10.6 10.5 10.8 0.3HL 10.6 10.0 11.4 10.2 10.0 8.4 9.1 10.0 1.0IUE 2.4 2.0 2.4 2.4 2.2 2.5 2.7 2.4 0.2

UEW 3.3 2.8 3.1 3.1 3.1 3.1 3.2 3.1 0.2SL 3.2 3.1 3.2 3.7 3.8 3.2 3.2 3.3 0.3EL 4.1 4.2 4.2 4.7 4.5 4.2 4.2 4.3 0.2FLL 6.2 5.3 5.8 5.8 6.2 5.2 6.0 5.8 0.4HAL 6.3 6.0 7.1 7.6 7.3 6.0 7.1 6.8 0.7ShL 11.8 10.1 11.5 11.3 11.5 10.2 11.2 11.1 0.7TL 11.9 10.0 11.6 11.2 11.6 10.1 11.3 11.1 0.8

FOL 11.9 10.2 11.5 11.4 11.5 10.3 11.5 11.2 0.7

* holotype and ** paratype of Bufo sulphureus Grandison and Daniel, 1964.

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Natural History Society. In this study, we examined theholotype and 18 paratypes (14 adult males - SVL 24.0-31.8 mm and 4 subadults - SVL 20.9-25.0 mm) depositedin BNHS and five paratypes in BMNH under the nomen'Bufo sulphureus'. In addition, recent collections of 'Bufo'koynayensis from the type locality were compared. Allavailable material of this species is consistent with theoriginal description.

ConclusionGhatophryne gen. nov. and Xanthophryne gen. nov. are bothendemic to the Western Ghats of India. Together with thearboreal toad genus Pedostibes (Western Ghats endemic;species from South-East Asia belong to another clade) andthe genus Adenomus (Sri Lanka endemic), they form theoldest lineages in the Adenominae clade. The recognitionof each of these lineages as distinct genera highlights theirearly diversification on the southern parts of the Indiansubcontinent [5].

Competing interestsThe authors declare that they have no competing interests.

Authors' contributionsAll authors contributed equally, and read and approvedthe final manuscript.

AcknowledgementsWe thank G. Ramakrishna and Kausik Deuti (ZSI), Asad Rahmani, J.C. Dan-iel (BNHS) and B. Clarke (NHM) for their support in our studies. Anil Zach-arya, Kehimkar, I. Agarwal and V. Patil provided valuable support in the field; I. Das provided valuable literature; I. Agarwal and Hemant Ogale made photographs of Xanthophryne available. The State Forest Departments of Kerala and Maharashtra kindly provided study permits to SDB and VG, and are greatly acknowledged. SDB is grateful to the Ministry of Environment

and Forest (MoEF) and University of Delhi (Scheme to strengthen R&D and Doctoral Research Program) for funds for the research. IVB is supported by a research grant from the Institute for the Promotion of Innovation through Science and Technology in Flanders (IWT-Vlaanderen). FB receives a postdoctoral fellowship from the Fonds voor Wetenschappelijk Onder-zoek-Vlaanderen (FWO). Financial support was provided FWO-Vlaan-deren (grants FWO 1.5.039.03N, FWO G.0056.03, FWO G.0307.04) and the Vrije Universiteit Brussel (grant VUB OZR834).

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Xanthophryne tigerinus eggs are laid in clutches in temporary puddlesFigure 4Xanthophryne tigerinus eggs are laid in clutches in temporary puddles.

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